JZAR Evidence-based practice

Journal of Zoo andAquarium Research 4(1)2016 OPEN ACCESS qiaet we i i hue idos Al wavelengths All indoors. housed is it when equivalent, artificial the or – heat and light solar of experience daily its by Reptile andAmphibian Working Group (RAWG). (BIAZA) Aquaria and Zoos of Association Irish and British the of Group Focus UV the by initiated was project The captivity. in kept amphibians and reptiles all for lighting UV suitable to guide a as used be can that document working a create to is project this of aim The non-existent. almost is gradients, UV satisfactory achieve to how and species, different for UV-B of 2014; Tapley et al. 2015). However, guidance as Johnson toand Carmel suitable2006; Rossi (e.g. levels recommended widely is The provision of UV lighting to captive reptiles and amphibians Introduction 2016 Published online:31January Accepted: 2016 15January Received: 9July2015 Article history: lighting UV lamps, UVrequirements, vivarium microhabitat design,UV-B, UVindex, Keywords: 7 6 5 4 3 2 1 Frances Baines selection ofUVlighting for reptiles andamphibiansin captivity How muchUV-B does my reptile need?TheUV-Tool, aguide to the Evidence-based practice *Correspondence: Frances Baines,UVGuideUK,Greenfield,SchoolLane,Govilon, Abergavenny NP79NT, UK;[email protected] Bristol Zoo Gardens,UK Durrell Wildlife Conservation Trust, Jersey Zoological Society ofLondon,UK Zoo,Marwell UK Chester Zoo, UK Reaseheath College,Nantwich, UK UV GuideUK,Abergavenny, UK Every aspect of the life of a reptile or amphibian is governed

1* , JoeChattell

2 , JamesDale (BIAZA) Reptile and Amphibian Working Group (RAWG) with contributions from zookeepers and from zookeepers with contributions herpetologists from theUKandabroad. Furtherinputiswelcome andencouraged. (RAWG) Group Working Amphibian and Aquaria Reptile and Zoos of (BIAZA) Association Irish and British the of members by compiled been has document UV-Tool. the of version current This the in different products 24 on reports are There enclosures. zoo larger in and vivaria in use for lamps appropriate of selection to guide a and products, lighting UV-B available commercially for maps gradient UV-index and reports test includes UV-Tooltherefore The used. be to lamps the of output and spectrum UV the environment.of Torequiresknowledge this do safeeffectiveplan tonextstepbut is ascertained,the captive the is UV within gradients amphibian or reptile of species any requirementof likelyUV the UV-index Once upon measurements. based zones) The possible. UV-ToolUV-Bfour‘zones’the of currentversionexposureof of (Ferguson each is then to species 254 assigns exposure UV-B of daily range natural of its estimation microhabitat, native its from inferred observed or behaviour in captivity. studies, Since field an animal’s from UV-B derived exposure is determined as by its behaviour species, within amphibian and reptile of behaviour basking UV-ToolThe wild. microhabitatexpandingcontainsthe databaseeditableand of requirementsan and thatseeks to address problem,this by considering rangethe experiencedUV of the by in species each document aworking The is or lighting. UV-Tool artificial amphibians, using gradients UV and satisfactory achieve reptiles to how for lighting UV of levels suitable to as non-existent almost is Guidance Abstract 3 , DanGarrick 4 , IriGill 5 , Matt Goetz iai D pre- vitamin to skin, the in sterol a (7DHC), 7-dehydrocholesterol of within thespectrumistherefore very important. provision its 2002); al. et Honkavaara 1993; al. et Fleishman 1974; Moehn 1974; Zueva and (Govardovskii items food and reptiles and amphibians, which use it in recognising conspecifics 400 nm). (320– UV-A fraction, longer-wavelength a and nm) (290–320 UV-B fraction, short-wavelength a of consists sunlight natural wavelength;by subdivided is It sunlight.component of normal a is Ultraviolet patterns. activity daily their and microhabitat these by their upon depend that utilised amounts in received are and be animals, may (UV) ultraviolet to infra-red from Short-wavelength UV-B (290–315nm) enables the conversion UV-A from around 350 nm is within the visual range of many 3 I si ti udros temperature-dependent a undergoes this skin In . 6 , TimSkelton 7 andMatt Swatman 3 42 A UV-B lighting guide for reptiles and amphibians

isomerisation into vitamin D3, which is metabolised by the liver 1988, 1989, 1991; Hertz et al 1994; Dickinson and Fa 1997) and and subsequently by the kidney into the vital endocrine hormone in some cases for UV photoregulation (Manning and Grigg 1997; calcitriol, controlling calcium metabolism. It is also metabolised into Ferguson et al. 2003; Karsten et al. 2009). The animal’s response calcitriol intracellularly throughout the body, where in mammals will determine its exposure within these gradients. Variation in it has been shown to perform multiple autocrine and paracrine behaviour creates enormous differences in UV exposure between functions, controlling transcription of as many as 2000 genes species, ranging from mid-day full-sun baskers to nocturnal and that influence functions as diverse as growth, insulin production crepuscular animals, which may receive the majority of their and the immune system (Hossein-nezhad and Holick 2013). ultraviolet exposure from small amounts of daylight reaching

Overproduction of vitamin D3 is prevented by the conversion of them in their diurnal retreats.

excess pre-vitamin D3 and vitamin D3 into inert photoproducts by The creation of similar superimposed heat, light and UV UV-B and short-wavelength UV-A (range 290–335nm), effectively gradients using UV-B-emitting lamps, often in combination making this natural process, in sunlight, self-limiting (MacLaughlin with other sources of heat and light, is possible because their et al. 1982; Webb et al. 1989). Although most research on vitamin irradiance is proportional to the distance from the lamp. The task

D3 has been carried out on mammals, studies conducted on other requires knowledge of (1) the range of irradiance appropriate for taxa indicate that vitamin D pathways are similar in most terrestrial the species and (2) the gradients created by individual lighting vertebrates (Holick et al. 1995; Bidmon and Stumpf 1996; Antwis products, which may be used individually or in combination to and Browne 2009). produce the desired effect. As well as its role in enabling and regulating cutaneous vitamin D synthesis, UV has direct effects upon skin, which include The range of irradiance appropriate for the species modulation of the cutaneous immune system, strengthening of Research on this topic is in its infancy, even with regard to skin barrier functions and increasing pigment formation. It also human beings. There is hardly any scientific data to back the stimulates production of beta endorphins, giving sunlight its recommendation of any particular level of UV-B for any particular ‘feel good’ factor, and induces nitric oxide production, which has species. Until very recently, no practical methods existed for localised protective effects (Juzeniene and Moan 2012). Solar UV recording ambient UV-B in the microhabitat of free-living reptiles is also an effective disinfectant (McGuigan et al. 2012) that can and amphibians. However, Ferguson et al. (2010) reported the UV destroy bacteria, fungi and viruses on the surface of the skin. exposure of 15 species of reptiles in the field during their daily Excessive exposure to UV must, however, be avoided. High and seasonal peak of activity, using the unitless UV index (UVI), doses and/or exposure to unnaturally short-wavelength UV from as measured with a Solarmeter 6.5 UV Index meter (Solartech artificial sources can result in eye and skin damage, reproductive Inc., Harrison Township, Michigan, USA), and demonstrated failure or even the death of amphibians (Blaustein and Belden that knowledge of the basking/daylight exposure habits of any 2003) and reptiles (Gardiner et al. 2009), and in mammals, can species enables a reasonable estimation of likely UV exposures to lead to the formation of skin cancers (Soehnge 1997). Squamous be made. They allocated species into four sun exposure groups cell carcinomas have been reported in captive reptiles but the or ‘zones’, which have since been designated ‘UV-B Zones’ or significance of their association with the use of artificial UV ‘Ferguson zones’ (Carmel and Johnson 2014; Ferguson et al. 2014). lighting is as yet undetermined (Duarte and Baines 2009; Hannon For each zone, a range of figures was given for the mean voluntary et al. 2011). UVI exposures calculated from all readings (Zone Range), and for Species vary widely in their basking behaviours or lack of them the maximum UVI in which the animals were encountered. The (Avery 1982; Tattersall et al. 2006; Michaels and Preziosi 2013), Ferguson zones are summarised in Table 1. their skin permeability to UV radiation (Porter 1967; Nietzke 1990) Any species can be assigned to one of the four zones based

and in their response to UV-B in terms of vitamin D3 production upon its basking behaviour. The authors suggest that a suitable UV (Carman et al. 2000). These behavioural and morphological gradient may then be provided in the captive animal’s environment characteristics optimise their UV exposure for vitamin D synthesis using these figures as a guide. Such a gradient should enable the and the other beneficial effects of sunlight, whilst simultaneously animal to self-regulate its exposure from zero (full shade) to the minimising the risk of UV damage, but these adaptations are only maximum indicated for that zone, which would be provided at the relevant for the solar irradiation they experience in their native animal’s closest access point to the lamp. microhabitat. Thus it would seem very important to match the solar UV spectrum as closely as possible, and to recreate the levels of irradiance found in this microhabitat, when providing reptiles and amphibians with artificial lighting. Table 1. The Ferguson zones, summarised from Ferguson et al. (2010). In nature the levels of UV irradiance at any one location vary Species are grouped into four zones according to their thermoregulatory behaviour and microhabitat preferences, with the UVB reference continuously, unlike the situation in a typical vivarium, in which a guidelines determined from average irradiance of randomly encountered UV-B-emitting lamp is either on or off. The greatest determinant individuals in the field. of irradiance is the solar altitude – the height of the sun in the sky – because at low solar altitudes the rays must pass through a Zone range Maximum thicker layer of atmosphere, which selectively absorbs and scatters Characteristics UVI UVI shorter wavelengths. Under clear skies, the solar UV-B levels rise Crepuscular or shade dweller, from zero at dawn, to a maximum around noon, then fall again Zone 1 0–0.7 0.6–1.4 thermal conformer to zero at sunset (e.g. Michaels and Preziosi 2013). Clouds scatter and absorb all wavelengths, and may greatly reduce irradiance. Partial sun/occasional basker, Zone 2 0.7–1.0 1.1–3.0 However, meteorological data cannot be representative of thermoregulator conditions within a microclimate. At any time of day, sunlight also interacts with features in the animal’s environment such as trees, Open or partial sun basker, Zone 3 1.0–2.6 2.9–7.4 rocks, plants and water, creating superimposed gradients of heat, thermoregulator light and UV extending from full sunlight into full shade. Reptiles Mid-day sun basker, and amphibians perceive these gradients and may use light Zone 4 2.6–3.5 4.5–9.5 intensity as a cue for thermoregulation (Sievert and Hutchison thermoregulator

Journal of Zoo and Aquarium Research 4(1) 2016 43 Baines et al.

The gradients created by individual lighting products information on the animal’s natural microhabitat and thermal The suitability of any light source is governed by two main features: requirements was added, to assist the keeper in choosing its quality (the spectrum) and quantity (the irradiance received by appropriate lamp combinations for creating a suitable lighting and the animal). The template for the ideal spectral power distribution heating gradient within the enclosure. The database included the is the solar spectrum, under which life evolved and to which all following information: life on the planet’s surface is adapted. Direct comparisons of lamp • Species (Latin name, common name) spectra with the solar spectrum are therefore required. • Biome (Major biome or Terrestrial Ecoregion as defined by With regard to quantity, the irradiance at any given distance Olson et al. (2001) and adopted by the World Wildlife Fund from a lamp is a function of the output of the lamp and the way (WWF 2015) the light is distributed, i.e. the shape of the beam. For example, • Ferguson zone a fluorescent tube that radiates a diffuse, relatively low level of • Photoperiod UV-B from its entire surface will produce a very different UV-B • Winter treatment, if any (cooling, brumation or gradient and basking opportunity than a mercury vapour spot hibernation) lamp that emits a very narrow beam of intense UV-B light only • Basking zone temperature (substrate surface a few centimetres across. The use of various lamp reflectors, temperature) shades or luminaires can also dramatically affect the shape of the • Daytime ambient (air) temperature (summer and winter) beam and the intensity of UV at any given distance. It is therefore • Night ambient (air) temperature (summer and winter) important to plot an iso-irradiance chart for each lamp, to assess • Microhabitat, including specialist requirements added as its effectiveness. However, in previous studies the irradiance from ‘comments’ UV-B lamps has usually been measured at standard distances from the lamp, regardless of the lamp type and the shape of its A selection of 24 widely available UV-B-emitting lighting beam (Gehrmann 1987; Gehrmann et al. 2004b; Lindgren 2004; products was fully tested by one of the authors (FB). The lamps Lindgren et al. 2008). were switched on for 15 hours per day until a total of 105 hours A hand-held broadband meter is a practical instrument was completed before testing, approximating the industry for measuring both solar UV irradiance in the field and lamp standard ‘burning-in’ period of 100 hours (IESNA 1999). irradiance indoors. However, different brands and models of All measurements were carried out with the lamps in simple broadband UV-B meters (range 280–320 nm) will have different fixtures, with no shades or reflectors, above a test bench, after a spectral responsivity. Unless they are specifically calibrated for 30-minute warm-up period. Recordings included: the spectral power distribution of a particular lamp, each meter • Spectrograms (Ocean Optics USB2000+ spectral radiometer may give a different reading from that lamp at any given distance with a UV-B compatible fibre-optic probe with cosine (Gehrmann et al. 2004a). In addition, only a very narrow band of adaptor: Ocean Optics Inc., Dunedin, FL 34698 USA) shorter wavelengths in the UV-B range (295–315 nm) contribute • UV Index (Solarmeter 6.5 UV Index meter: Solartech Inc.,

to vitamin D3 synthesis; measurements including irradiance from Harrison Township, MI 48045 USA) longer wavelengths may be misleading as to the effectiveness of • Total UV-B: 280–320nm (Solarmeter 6.2 broadband UVB a lamp. meter: Solartech Inc., Harrison Township, MI 48045 USA) Unlike broadband UV-B meters, which respond to the entire • UV-C (Solarmeter 8.0 broadband UVC meter: Solartech range of UV-B wavelengths, the Solarmeter 6.5 UV Index Inc., Harrison Township, MI 48045 USA) meter (Solartech Inc., Harrison Township, Michigan, USA) used • Visible light output (SkyTronic LX101 model 600.620 digital by Ferguson et al. (2010) has strong filtration of the longer lux meter: SkyTronic B. V., Overijssel, Netherlands) wavelengths, resulting in a spectral responsivity with a 96% • Electrical consumption (Prodigit power monitor model

overlap to the CIE pre-vitamin D3 spectrum (CIE 2006) from 290 to 2000M-UK: Prodigit Electronics, New Taipei City, Taiwan) 400 nm (S. Wunderlich, pers. comm.). This enables a reasonable estimate of the vitamin D-synthesising potential of sunlight and For those lamps emitting UV-B in appropriate wavelengths for

any artificial source. The readings are displayed in the unitless UV vitamin D3 synthesis, as indicated by their spectral analysis, an iso- index, which is beneficial for interpretation as it is a well known irradiance chart mapping the UV index gradient was constructed measurement of ‘sun strength’ as determined by human erythema, according to a method described previously (Baines 2015). The which has a similar, but not identical, action spectrum (CIE 1998) ability of each lighting product to provide irradiances within the UV

to the pre-vitamin D3 spectrum. The Solarmeter 6.5’s spectral index ranges appropriate to each Ferguson zone was documented response falls about halfway between the two (Schmalwieser et and guidelines drafted regarding methods of lamp choice. al. 2006). When its UVI measurements were compared with data The species database, lamp test results and guidelines were provided by a Bentham spectrometer, a very accurate sensor used compiled into a draft Excel document. This was distributed to for UV measurements, deviations of only ±5% were found, which the wider BIAZA RAWG community and to a small number of are within the range commonly expected for scientific instruments herpetologists and private keepers with specialist knowledge. (de Paula Corrêa et al. 2010). This meter is therefore suitable for All recipients of the draft document were requested to submit measuring the irradiance from sunlight and from a lamp at specific reviews of the UV-Tool and data for additional species held in distances, and for plotting the shape of the lamp’s beam, to create their collections, including references to their source material an iso-irradiance chart. where appropriate. The first draft was distributed in December 2012, listing 190 species from the five zoological collections to Methods which the co-authors were affiliated. Between January 2013 and October 2015 contributions were received from a further nine A database was compiled of basic information on each species of institutions and ten individual contributors, bringing the total up reptile and amphibian held by the authors’ current institutions. to 254 species of reptiles and amphibians. This is still a working Each species was assigned to a Ferguson zone based on an document. The database has been updated at regular intervals, assessment of its basking behaviour, derived from published or and is currently in its tenth edition, available for download from personal studies made in the field if possible, but if not, from the Internet (BIAZA RAWG 2015). New reviews, corrections and observations of the animal’s behaviour in captivity. Further submissions are welcomed.

44 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians

Table 2. Assessment of 24 lamps used in reptile husbandry. Operating ranges also respect safe minimum distances. Fluorescent lamps emitting less than UVI 0.5 at 15cm are not considered to be suitable as the sole source of UVB even for Zone 1 species.

Ferguson zones which can be covered using the lamp (depending upon distance) Zone 1 Zone 2 Zone 3 Zone 4

Sample in Date sample using shade using shade using sunbeam using sunbeam Company name Brand name this report purchased method method method method

Fluorescent tubes A) T8 (1" diameter) tubes Arcadia Natural Sunlight Lamp 2% UVB 60cm 18W 2008 with reflector Arcadia D3 Reptile Lamp 6% UVB 60cm 18W 2008  with reflector Arcadia D3+ Reptile Lamp 12% UVB 60cm 18W 2008   with reflector Narva BioVital T8 60cm 18W 2009  ZooMed Reptisun 2.0/ Naturesun 60cm 18W 2008  ZooMed Reptisun 5.0/ IguanaLight 60cm 18W 2005   ZooMed Reptisun 10.0 60cm 18W 2011   with reflector B) T5 (16mm diameter) tubes Arcadia T5 D3 Reptile Lamp 6% UVB 55cm 24W 2011   with reflector with reflector Arcadia T5 D3+ Reptile Lamp 12% UVB 55cm 24W 2011     ZooMed Reptisun 5.0 UVB T5-HO 55cm 24W 2012   with reflector with reflector ZooMed Reptisun 10.0 UVB T5-HO 55cm 24W 2012     Mercury vapour lamps Arcadia D3 Basking Lamp 100W 2012   Arcadia D3 Basking Lamp 160W 2012   ExoTerra Solar Glo 125W 2012-2013   ? ExoTerra Solar Glo 160W 2012-2013   MegaRay PetCare Mega-Ray 100W 2014     MegaRay PetCare Mega-Ray 160W 2014     Osram Ultravitalux 300W 2005-2011     ZooMed Powersun 100W 2012-2013    ZooMed Powersun 160W 2012-2013     Metal halide lamps Color Arc manufactured prior to Iwasaki EYE 2011 150W 2009-2010   ? Iwasaki EYE Color Arc manufactured after 2011 150W 2009-2010  Lucky Reptile Bright Sun UV Desert 35W 2012    

Lucky Reptile Bright Sun UV Desert 50W 2008  

Results UV-B lamp test results Table 2 lists the lamps that were included in the trial, and Species database summarises their ability to provide irradiances within the UV index The entries to date (254 species) are listed in full in the Appendix. ranges appropriate to each Ferguson zone, at practical distances The contributors for each species and their recommended reading beneath the lamp. Figure 1A–C graphs the UVI irradiances of and reference lists are not included owing to space limitations, individual lamps at increasing distances from the surface of but are present in the UV-Tool Excel working document available the lamp, with the UV index meter positioned perpendicular to online (BIAZA RAWG 2015). Further contributions are still being the lamp, directly beneath its central point. Figures 2 and 3 are sought, and the BIAZA RAWG Focus Group intends to edit and examples of iso-irradiance charts and spectra for four distinct expand the database as more information becomes available. types of UV-B-emitting lamp: a standard-output T8 (25 mm

Journal of Zoo and Aquarium Research 4(1) 2016 45 Baines et al.

Figure 1. UV Index irradiance recordings. (A) UVB-emitting fluorescent tubes (T8 and T5 versions); (B) mercury vapour lamps; (C) metal halide lamps.

diameter) fluorescent tube, a mercury vapour lamp, a metal halide Discussion lamp and a T5 (16 mm diameter) High-Output (T5-HO) fluorescent tube fitted with an aluminium reflector. Each of the full lamp test Lamp test results results for all 24 lamps are accessible from links on the same The UV output of lamps sold for use with reptiles and amphibians website page from which the Excel working document may be varies enormously, not just from different types of lamp, but also downloaded (BIAZA RAWG 2015), as well as from links within the from different brands with similar specifications. Although only UV-Tool itself. New lamp test results will be added to this website, one lamp from each brand was tested in this trial, previous tests and their links will be added to the working document, as they (FB, unpublished data) have shown that considerable differences become available. may exist between individual lamps of the same brand and

46 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians

A B

C D

Figure 2. Full ultraviolet and visible light (UV-VIS) spectrum of samples of four types of UVB lamp. A mid-day solar spectrum with the sun close to the zenith (Solar altitude 85.4°) is overlaid onto each chart - but note the different irradiance scales. This enables comparison of the spectral power distribution of the lamp with that of natural sunlight, which has a completely continuous spectrum from a threshold around 295nm. (A) UVB-emitting fluorescent tube (T8 version): ZooMed Reptisun 10.0 UVB 18watt T8 fluorescent tube. Distance 10cm. (B) Mercury vapour lamp: ZooMed Powersun 160watt lamp. Distance 30cm. (C) Metal halide lamp: Lucky Reptile Bright Sun Desert UV 50watt lamp. Distance 30cm. (D) UVB-emitting fluorescent tube (T5 version): Arcadia T5- HO D3+ 12%UVB 24watt T5 fluorescent tube in aluminium reflector. Distance 10cm.

specifications. This may be due to small differences in manufacture D3 synthesis, although the so-called ‘full spectrum’ fluorescent such as internal positioning of lamp elements, thickness of glass or tubes, Narva Biovital (Narva Lichtquellen GmbH, Brand-Erbisdorf, coatings, etc., but the UV-B output may also vary with external Germany), ZooMed NatureSun (ZooMed Laboratories Inc., San factors such as fluctuations in the voltage of the electrical supply Luis Obispo, USA), and the Iwasaki EYE Color Arc metal halide lamp and the ambient temperature. UV-B output also decays with manufactured after 2011 (Iwasaki Electric Co. Ltd., Tokyo, Japan), use, primarily due to solarisation of the glass envelope under UV emit insignificant amounts except at extremely close range. bombardment, but also due to chemical changes in phosphors or halide mixtures or the blackening of glass from sputtering from Iso-irradiance charts ageing electrodes. Ideally, lamp output should be monitored The iso-irradiance charts enable comparison of the UV gradients regularly. Most products decay only slowly, however, after the between different lamps and reveal important differences in initial ‘burning-in’ period. Included in the full lamp test results are the surface area beneath the lamps that receives any specified measurements taken from seven of the UV-emitting fluorescent irradiance. For example, as indicated in Table 2, both the Arcadia tubes from Arcadia (Arcadia Products plc., Redhill, UK) and T5 D3+ Reptile Lamp 12% UV-B fluorescent tube (Arcadia Products ZooMed (ZooMed Laboratories Inc., San Luis Obispo, USA), each plc., Redhill, UK) and the Lucky Reptile Bright Sun Desert 50watt lamp representing a different brand, put into use for at least a full metal halide lamp (Import Export Peter Hoch GmbH, Waldkirch, year (4000 hours of use at 10–12 hours per day). After burning-in Germany) are able to produce a gradient suitable for a Zone 2 for 105 hours, the mean reduction in UVI, from new, was 12.6% animal at a safe distance. However, the iso-irradiance charts (range 6–23%). At the end of 4000 hours the mean reduction in for these lamps indicate that the fluorescent tube fitted with a UVI, from new, was only 39.9% (range 30–48%). These results reflector provides a UV index range between 0.5 and 1.0 across suggest that some brands may not need replacement for at least an area over 130 cm in diameter at a distance of 85 cm (Figure one year. Not all products have similar longevity. For example, one 3D), whereas the same zone of irradiance under the metal halide brand sold by a different manufacturer showed a reduction in UVI lamp is achieved at 45 cm, but the footprint is less than 25 cm in of 64% from new after only 1000 hours’ use – about three months diameter (Figure 3C). The practical uses for these two lamps will at 10–12 hours per day. This product was therefore rendered therefore be very different. Effective UV coverage needs to be at ineffective at any practical distance after only three months (FB, least as wide as the whole body of the animal. unpublished data). Mercury vapour and metal halide lamps emit significant infrared Spectral analysis reveals that none of the lamps in this trial radiation as well as UV and visible light. When creating a thermal emit harmful non-solar UV-B radiation (<290 nm). All of the gradient, just as with a UV gradient, the whole of the animal’s lamps emit at least some UV-B in the range required for vitamin body must fit within the optimum upper temperature zone. For

Journal of Zoo and Aquarium Research 4(1) 2016 47 Baines et al.

Figure 3. UV Index iso-irradiance charts for samples of four types of UVB lamp. These charts are for the lamps with spectra illustrated in Figure 2. (A) UVB- emitting fluorescent tube (T8): ZooMed Reptisun 10.0 UVB 18watt T8 tube. (B) Mercury vapour lamp: ZooMed Powersun 160watt lamp. (C) Metal halide lamp: Lucky Reptile Bright Sun Desert UV 50watt lamp. (D) UVB-emitting fluorescent tube (T5): Arcadia T5-HO D3+ 12%UVB 24watt T5 fluorescent tube in aluminium reflector.

basking species, this means creation of a basking area in which 2. ‘Max UVI recorded’ refers to the highest UVI that the appropriate UV, visible light and infrared radiation cover the entire reptiles from each zone were found to occupy in this study. body of the animal. Lamps with wide flood-type beams or multiple Obviously this figure might reflect a ‘one-off’ exposure – a lamps above the basking zone are often necessary. single reptile found out in mid-day sun – but it gives an When lamps are installed in enclosures, any shades and estimate of the maximum levels this type of animal might reflectors, mesh guards, even nearby objects such as branches encounter naturally. This might be considered as a guide and foliage will affect light and UV distribution. Iso-irradiance as to the upper acceptable limit for the UV gradient to be charts are no substitute for in-situ measurements; they are merely provided in captivity. guides to aid lamp selection. We suggest that a suitable UV gradient, chosen to match the Using the Ferguson zones zone to which the reptile or amphibian is allocated, may then be Figure 4 summarises the zone ranges recorded by Ferguson et al. provided in the captive animal’s environment, enabling the animal (2010) and illustrates the way in which we propose they might be to self-regulate its exposure. A full range of UV levels may be used to create suitable UV gradients for any species based upon its provided, from zero (full shade) to the maximum suggested by the thermoregulation behaviour. zone assessment (at the closest point possible between the animal Ferguson et al. (2010) provide two sets of figures: and the lamp). We have used the information from the Ferguson 1. ‘Zone ranges’: all the UVI readings for the microhabitats at et al. (2010) study as a basis for our suggestion that there are two the time and place the reptiles were found were averaged. ways of supplying UV to reptiles and amphibians kept indoors in For example, the average exposure of crepuscular or shade captivity. dwelling species fell in the range between UVI 0 and 0.7, the ‘partial sun or occasional baskers’ were in a range from ‘Shade’ and ‘sunbeam’ methods 0.7 to 1.0, and so on. This figure might be considered a The ‘shade method’ provides low-level background UV over a suitable ‘mid-background’ level of UV for the species in large proportion of the animal’s enclosure using the zone ranges question. as a guide to appropriate ambient UV, with a gradient from the

48 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians

Figure 4. UV index estimates based upon the Ferguson zones. Columns 1 to 5 of the table identify the characteristics of each zone as presented by Ferguson et al. (2010). The original 15 species of reptiles studied in their natural habitat in Jamaica and south and west USA are shown in column 5. In the column 6 are examples of species commonly held in captivity, assigned to Ferguson zones based upon their known basking behaviour. Arrows link animals from each zone to either shade or sunbeam methods of UV provision as proposed in the BIAZA UV-Tool (2012) and indicate typical lamp types suggested for each method.

highest zone range value close to the lamp, to zero in the shade. the maximum permitted for each zone, with the exception of zone This would seem to be the method of choice for shade-dwelling 4. Although some zone 4 reptiles have been observed basking at animals and occasional baskers, i.e., those in zones 1 and 2. Most UVI 9.5 or above (Ferguson et al. 2010, 2014), even these spend amphibians, snakes and crepuscular lizards have been allocated to the majority of their basking time in the early morning and late these zones. Fluorescent T8 (26 mm diameter) UV-B tubes create afternoon, when levels are around UVI 3.0–5.0. It follows that low levels of UV irradiance, similar to those found in outdoor shade the most appropriate levels for zone 4 animals, too, will be in on a sunny day, over a relatively large area close to the tube, with this range. We suggest that for safety, UVI 7.0–8.0 should be a gradient to zero at greater distances from the lamp. They would considered the absolute maximum UVI at reptile level for zone 4 therefore appear to be particularly suitable for the shade method reptiles under artificial sources of UV-B, since the UV spectrum in small enclosures and vivaria, where the maximum UVI required from artificial lighting is not the same as from natural sunlight. would be no higher than UVI 0.7–1.0. In larger enclosures, high If keepers do not have access to a UV index meter, the iso- output T5 (T5-HO) (16 mm diameter) UV-B fluorescent tubes may irradiance charts and irradiance tables to which the UV-Tool be used, as these can be positioned further from the animals, to is linked may be used to identify suitable distances at which achieve the same low UVI at animal level. appropriate levels for both ‘shade’ and ‘sunbeam’ methods are The ‘sunbeam method’ is designed to provide a higher level achieved by different lamps. of UV for species known to bask in direct sunlight. The aim is to provide UV levels in the basking area that are similar to those Special considerations: nocturnal species experienced by a wild animal in direct sunlight in its natural Traditionally, it has been assumed that nocturnal and crepuscular habitat during a typical early to mid-morning basking period. This species do not require UV lighting because their lifestyle precludes

is the time when most basking species absorb solar radiation for exposure to daylight, and/or they obtain all the vitamin D3 they long periods. In the tropics and sub-tropics, in open sunlight on require from their diet. Although carnivores may obtain sufficient

clear days between 8.30am and 9.30am local time, the UV index vitamin D3 from the bodies of their prey, the natural diets of is typically in the range UVI 3.0–5.0 (FB, unpublished data). This insectivores are unlikely to provide any significant amounts of the higher level needs to be restricted to the basking zone (simulating vitamin (Finke and Oonincx 2014), making cutaneous synthesis a patch of sunlight) with a gradient to zero into shade. This method the most likely primary source. would seem appropriate for animals in zones 3 and 4, many of More than 60 years ago, reports were collected of supposedly which are diurnal reptiles, and for some partial sun/occasional nocturnal reptiles experiencing at least some exposure to baskers from zone 2. Some mercury vapour lamps, metal halide daylight, either by occasional daytime forays or by incidental UV-B lamps and high output T5 (T5-HO) UV-B fluorescent tubes exposure to light in their sleeping places (Brattstrom 1952). House (16 mm diameter) can produce much higher levels of UV-B than geckos, Hemidactylus frenatus and H. turcicus, are often seen T8 fluorescent tubes, up to levels typical of natural sunlight. These in daylight around dusk and dawn (FB, pers. obs.) and Tarentola lamps can be positioned to irradiate a brightly illuminated basking mauretanica can regularly be seen basking in the sun for periods zone with appropriate levels of UV-B for the entire photoperiod, throughout the day (MG, pers. obs.). Without evidence from 24- so that suitable UV exposure occurs whenever the animal chooses hour observational field studies, it cannot be assumed that any to bask. We suggest that ‘Max UVI recorded’ should be a guide to nocturnal species receives no sunlight at all. Many snakes, such as

Journal of Zoo and Aquarium Research 4(1) 2016 49 Baines et al.

the black ratsnake (Pantherophis obsoletus) vary their diel patterns 2009). They are therefore likely to need much reduced exposure

of activity depending upon ambient temperatures, increasing levels. Fortunately adequate vitamin D3 synthesis should still diurnal activity in the cooler months (Sperry et al. 2013). be possible despite lower UV exposure, since reduced melanin It has been speculated that crepuscular species may synthesise pigment allows more UV-B to enter the epidermal cells.

vitamin D3 by emerging into sunlight at dusk and dawn. However, when the sun is close to the horizon, the atmosphere filters out Ontogenetic changes

almost all the UV-B wavelengths required for vitamin D3 synthesis; Consideration should also be given to any ontogenetic changes species which can benefit from such low levels of UV need skin in microhabitat and/or behaviour when allocating species to with very high UV transmission. Some nocturnal geckos, for Ferguson zones. Amphibians with both larval and adult life stages example, fit into this category. Short wavelength UV-B has been are obvious examples, but juvenile reptiles of many species also live shown to be transmitted through the full thickness of skin of the more cryptic lifestyles than the adults, inhabiting more sheltered nocturnal gecko Coleonyx variegatus to a depth of 1.2 to 1.9 mm, microhabitats with relatively less ambient UV. A well-known in stark comparison with diurnal species such as the desert lizard example of this is the Komodo dragon (Varanus komodoensis); Uta stansburiana, in which transmission was restricted to between juveniles are arboreal, whereas adults are ground-dwellers 0.3 and 0.9 mm (Porter 1967). In the same study, Porter found foraging across open savanna as well as in woodlands (Auffenberg that the skin transmission of seven species of snake reflected 1981). More fieldwork is needed to identify differences in the UV their behaviour, such that the highest transmission was seen in exposure of immature animals, to determine whether they need a the most completely nocturnal species, and the lowest in diurnal different Ferguson zone allocation from that of adults. Estimating species, with crepuscular snakes in between. This suggests one juvenile requirements was outside the remit of this project, but

way in which low levels of UV-B may enable adequate vitamin D3 these might usefully be added to the UV-Tool in the future. synthesis in nocturnal species. Carman et al. (2000) demonstrated that the skin of the nocturnal house gecko Hemidactylus turcicus General cautions

can synthesise vitamin D3 eight times more efficiently than skin In applying these guidelines to the provision of UV lighting, some from the diurnal desert lizard Sceloporus olivaceous – suggesting general cautions must be emphasised. that this is an adaptation either to lower levels of available Firstly, this is a very simplistic assessment, with very wide ultraviolet light in its microhabitat, or to very short exposure to interpretations possible. This is intentional; the concept is higher levels, during brief day-time emergences from shelter. designed to enable creation of wide, safe UV gradients combined Leopard geckos (Eublepharis macularius) synthesised vitamin with heat and light gradients, enabling reptiles and amphibians

D3 when exposed to low-level UV-B; 25-hydroxyvitamin D3 levels to photoregulate and thermoregulate simultaneously, throughout in exposed animals were 3.2 times higher than controls receiving the day. This requires the sources of UV, visible light and infrared only dietary supplementation (Wangen et al. 2013). Crepuscular radiation to be positioned close together, simulating sunlight, snakes such as the corn snake, Elaphe guttata, have also been and creating a basking zone at least as large as the whole body

shown to synthesise vitamin D3 in the skin when exposed to low of the animal. Multiple lamps may be required in some cases; levels of UV-B from fluorescent lamps (Acierno et al. 2008). the effects are additive for all wavelengths, so overlapping beams Mid-day UV-B filtering into the daylight sleeping places of must be used with caution. It also requires provision of adequate nocturnal animals may also be sufficient to enable adequate space and shelter, away from the lamps, for suitable gradients cutaneous synthesis. As far as we are aware, no published field to form. Provision of shade is vital for all species, regardless of studies exist recording the ambient UV-B in the daytime location their Ferguson zone. Even zone 4 reptiles must have a UV gradient of inactive nocturnal animals. However, UVI meter readings falling to zero in shelters away from the light. All guidelines to date between UVI 0.1 and 1.2 have been recorded beside leaf-tailed are still very experimental; the exact UV requirements of reptiles geckos (Uroplatus sp.) sleeping in daylight against tree trunks in and amphibians are still largely unknown, and it is vital to monitor Madagascar (L. Warren, pers. comm.) the animals’ responses and record results.

The vitamin D3 requirement of some nocturnal species may be Secondly, basking temperatures and ambient temperatures low; passive absorption of dietary calcium by vitamin D-deprived must be suitable, to ensure basking behaviours – and therefore leopard geckos, for example, appears to be effective enough to UV exposure times – are natural, neither abnormally short nor prevent metabolic bone disease (Allen et al. 1996). However, the prolonged.

paracrine and autocrine functions of vitamin D3 are independent Thirdly, lamps should always be positioned above the animal, from calcium metabolism; more research is needed to assess the so the shape of the head, and upper eyelids and eyebrow ridges full effects of vitamin D deficiency. when present, shade the eyes from the direct light. To summarise, some nocturnal animals clearly do have the Fourthly, all lamps present an electrical risk, and many also

ability to synthesise vitamin D3 in their skin, and this would occur present the risk of thermal burns and UV burns if the animal naturally whenever they were exposed to daylight. So there would can approach too closely. All bulbs should be inaccessible to the seem to be no reason to withhold provision of full spectrum animals; wire guards may be necessary. Wide wire mesh should lighting, provided that they are able to spend the daylight hours in be chosen where possible, to maximise light and UV transmission an appropriate retreat, with access to a UV-B component suitable (Burger et al. 2007). for a shade-dwelling or crepuscular species (i.e. Ferguson zone 1). Finally, ordinary glass or plastics must not be placed anywhere between the lamp and the animal, as these normally block Hypopigmentation transmission of all UV-B. Some high-transmission glass and Extra consideration is required when planning lighting for albino specialised UV-transmitting acrylics will, however, allow a certain and hypomelanistic specimens of any species, regardless of the proportion through, although even these materials selectively zone allocation of that species. Melanin strongly absorbs UV block shorter UV wavelengths. Spectral analysis conducted by radiation. A lack of skin and eye pigmentation therefore increases one of the authors (FB, unpublished data) indicated that 3 mm the transmission of radiation into the body (Solano 2014). Such UV-transmitting acrylic (Clear Sunbed Grade UV-T Perspex Acrylic animals are often popularly reported to be more sensitive to UV Sheet: Bay Plastics Ltd., North Shields, UK) permitted 80.9% and visible light (e.g. Dell’Amore 2007), and may be at increased transmission of UV-B at 300 nm. UV-transmitting twin-wall acrylic risk of UV-induced skin damage and cancer (Duarte and Baines roofing panels (Plexiglas Alltop SDP16: Evonik Industries AG,

50 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians

Essen, Germany) permitted 58.8% transmission at 300 nm. For Trust (Matt Goetz and Christopher Pye); Hadlow College, Kent comparison, a 4 mm sheet of high-transmission, low-iron glass (John Pemberton); Living Rainforest (Lisa Clifforde and Rob Ward); (Planibel Clearvision Glass: AGC Glass Europe, Louvain-La-Neuve, Marwell Wildlife (Dan Garrick); Newquay Zoo (Dan Garrick); Belgium) transmitted 16.9% of UV-B at 300 nm, compared to only Sparsholt College (Steve Nash); Wildlife and Wetlands Trust (Jay 0.4% transmission through ordinary 4 mm window glass. Redbond); ZSL London Zoo (Iri Gill, Ben Tapley and Sebastian Grant) and members of the Three Counties Tortoise Group. Guest Summary contributors: Andy Beveridge; Chris Davis; Gary Ferguson; Greg Fyfe; Jeffrey Lambert; Christopher Michaels; James Miller; Roman Very few field studies have been conducted on the natural UV Muryn; Jim Pether and Terry Thatcher. exposure of reptiles and amphibians. However, an estimation of a suitable UV range for any species may be made using knowledge References of its typical basking behaviour and its microhabitat. In indoor Acierno M.J, Mitchell M.A., Zachariah T.T., Roundtree M.K., Kirchgessner enclosures, careful positioning of UV lamps enables creation of a M.S., Sanchez-Migallon Guzman D. (2008) Effects of ultraviolet UV gradient within this range, which can be incorporated into full radiation on plasma 25-hydroxyvitamin D3 concentrations in corn snakes (Elaphe guttata). American Journal of Veterinary Research 69: spectrum lighting to simulate sunlight. 294–297. The BIAZA RAWG UV-Tool is a working document in which species Allen M.E., Oftedal O.T., Horst R.L. (1996) Remarkable differences in the are allocated to UVI ranges (Ferguson zones) according to their response to dietary vitamin D among species of reptiles and primates: basking behaviour. Information is also provided regarding suitable Is ultraviolet B light essential? In: Holick M.F., Jung E.G. (eds). Biologic temperature gradients, photoperiod and microhabitat, to assist Effects of Light 1995. Berlin: Walter de Gruyter, 13–30. Antwis R., Browne R. (2009) Ultraviolet radiation and vitamin D in construction of the photo-microhabitat. UV-B lamps vary widely 3 in output and beam characteristics, but links to lamp test results amphibian health, behaviour, diet and conservation. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology are available in the UV-Tool. Lamp choice will depend primarily on 154: 184–190. the Ferguson zone of the animal, which determines the required Auffenberg W. (1981) The Behavioral Ecology of the Komodo Monitor. UV gradient, and the size of the enclosure, which determines the Gainesville: University Press of Florida. distance at which the lamp can be placed. Final positioning of the Avery R.A. (1982) Field studies of body temperatures and thermoregulation. lamp (or lamps) is determined by using a UV index meter; if no In: Gans C., Pough, F.H. (eds). Biology of the Reptilia 12, Physiology C. meter is available, the charts and figures published in the test Physiological Ecology. London: Academic Press, 93–166. results may be helpful if the same lamps are being used. Baines F.M. (2015) Make yourself an iso-irradiance chart. A simple guide to mapping a UV index gradient. http://www.uvguide.co.uk/ Since this is a working document, we encourage submission of makingspreadcharts.htm (accessed 10 October 2015). new species data to the database, and updates of lamp test results BIAZA RAWG (2015) British and Irish Association of Zoos and Aquaria are planned. Reptile and Amphibian Working Group UV-TOOL PROJECT. http:// www.uvguide.co.uk/BIAZA-RAWG-UV-Tool.htm (accessed 10 October Definitions 2015). Bidmon H.J., Stumpf W.E. (1996) Vitamin D target systems in the brain of Irradiance is the radiant power received by a surface per unit area. the green lizard Anolis carolinensis. Anatomy and Embryology 193: 145–160. The units are microwatts per square centimetre (µW/cm²). Blaustein A.R., Belden, L.K. (2003) Amphibian defenses against ultraviolet-B radiation. Evolution & Development 5: 89–97. Illuminance is the total luminous flux received by a surface per Brattstrom, B.H. (1952). Diurnal activities of a nocturnal animal. unit area. This is a measure of the apparent brightness of an Herpetologica 8: 61–63. illuminated area to the human eye. It is calculated from the product Burger R.M., Gehrmann W.H., Ferguson G.W. (2007) Evaluation of UVB of the spectral irradiance (µW/cm² per nanometre of wavelength) reduction by materials commonly used in reptile husbandry. Zoo Biology 26: 417–423. with the human luminosity function, which represents the eye’s Carman E.N., Ferguson G.W., Gehrmann W.H., Chen T.C., Holick M.F. (2000) response to different wavelengths. This weighting is required Photobiosynthetic opportunity and ability for UV-B generated vitamin because human brightness perception is wavelength-dependent. D synthesis in free-living house geckos (Hemidactylus turcicus) and The unit is the lux. Since animal eyes have different spectral Texas spiny lizards (Sceloporus olivaceous). Copeia 2000: 245–250. sensitivities, it is only a crude estimate of the brightness perceived Carmel B., Johnson R. (2014) A Guide to Health and Disease in Reptiles & by any non-human species, but equivalent luminosity functions Amphibians. Burleigh, Australia: Reptile Publications. for reptile and amphibian species are lacking. CIE (1998) Erythema Reference Action Spectrum and Standard Erythema Dose. Vienna, Austria: Commission Internationale de l’Eclairage (International Commission on Illumination). Publication CIE The UV index (WHO 2002) is an international standard measurement S007E-1998. of the intensity of human erythemally-active (sunburn-producing) CIE (2006) Action Spectrum for the Production of Previtamin D in Human UV radiation. It is calculated from the product of the spectral Skin. Vienna, Austria: Commission Internationale de l’Eclairage irradiance (µW/cm² per nanometre of wavelength) and the human (International Commission on Illumination). Publication CIE 174– erythemal action spectrum across the range of UV wavelengths. 2006. This weighting is required because shorter UV wavelengths are de Paula Corrêa M., Godin-Beekmann S., Haeffelin M., Brogniez C., Verschaeve F., Saiag P., Pazmiño A., Mahé E. (2010) Comparison much more damaging than longer wavelengths. The UV index is between UV index measurements performed by research-grade and unitless. consumer-products instruments. Photochemical & Photobiological Sciences 9: 459–463. Acknowledgements Dell’Amore, C. (2007) Albino alligator makes zoo debut. http://news. We wish to thank the following contributors to the species nationalgeographic.com/news/2007/05/070514-white-gator.html microhabitat assessments. Zoological institutions and organisations: (accessed 10 October 2015). Birmingham Wildlife Conservation Park (Adam Radovanovic); Blue Dickinson H.C., Fa J.E. (1997) Ultraviolet light and heat source selection in captive spiny-tailed iguanas (Oplurus cuvieri). Zoo Biology 16: 391– Planet Aquarium (Joe Chattell); Bristol Zoo Gardens (Tim Skelton 401. and Adam Davis); Chessington World of Adventures (Keith Russell Duarte A.R., Baines F.M. (2009) Squamous cell carcinoma in a leopard and Rea Walshe); Chester Zoo (Matt Swatman and James Dale); gecko. Exotic DVM 11: 19–22. 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effective vitamin3 D -producing ultraviolet B radiation using Solartech’s WWF (2015) Terrestrial ecoregions. Washington DC: World Wildlife Solarmeter 6.4 handheld, UVB radiometer. Bulletin of the Chicago Fund. http://www.worldwildlife.org/biome-categories/terrestrial- Herpetological Society 43(4): 57–62. ecoregions (accessed 10 October 2015).

52 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians I I I J D IJ Microhabitat 6–9 different) Winter (if Winter temperature (°C) >20 Night – ambient (air) 24–26 12–15 24–25 24–26 Summer 12–17 23–25 different) Winter (if Winter 25 temperature (°C) Day – ambient (air) 13–20 25–30 26–30 25–30 25–30 Summer 30 35 30–35 31–36 30–35 30–32 Basking zone – Substrate surface temperature (°C) (if any) Hibernation Winter treatment Winter 13:11 13:11 13:11 14:10 12:12 12:12 Photoperiod Appendix: Microhabitat assessment Appendix: Microhabitat 3 2 2–3 2–3 2–3 2–3 Zone Ferguson Ferguson 1 4 1 1 1 1 Biome 01 Tropical and Subtropical Moist Broadleaf Forests; 02 Tropical and Subtropical Dry Broadleaf Forests; 03 Tropical and Subtropical Coniferous Forests; 04 Temperate Broadleaf and Mixed Forests; Forests; and Mixed Broadleaf 04 Temperate Forests; Coniferous and Subtropical 03 Tropical Forests; Dry Broadleaf and Subtropical 02 Tropical Forests; Broadleaf Moist and Subtropical 01 Tropical Common Name Spectacled Caiman Tuatara Cook Strait Philippine crocodile Morelet’s Crocodile Morelet’s Dwarf Crocodile Cuvier's Dwarf Caiman The animal undertakes preparation andgoes into atorpid forstate anextended period - duration in months.Physiological changes occur within the animal. Co-incident with winter and seen mostly in animals of The animal becomes torpid for a period of days or weeks. Co-incident with hotter weather. with hotter Co-incident or weeks. a period of days for torpid The animal becomes The animal becomes torpid for a period which may last weeks. Co-incident with winter. Co-incident weeks. last a period which may for torpid The animal becomes The temperature is reduced for a period, usually co-incident with winter. The animal reduces activity and feeding may cease, but it does not necessarily go into an extended, torpid state. torpid extended, an go into cease, but it does not necessarily feeding may activity and The animal reduces with winter. a period, usually co-incident for is reduced The temperature Key Biome biomes: WWF major terrestrial 10 and Savannas; and Shrublands; 09 Flooded Grasslands Savannas Grasslands, and Shrublands; 08 Temperate Savannas Grasslands, and Subtropical 07 Tropical Taiga; Forests/ 06 Boreal Forests; Coniferous 05 Temperate Shrublands; 14 Mangroves. and Scrub; 13 Deserts Xeric Woodlands Forests, 12 Mediterranean and Shrublands; 11 Tundra; Grasslands Montane behaviour Thermoregulatory zones: Ferguson sun basker. 4 - mid-day 3 - open or partial sun basker; basker; 2 - partial sun/ occasional or shade dweller; 1 - crepuscular treatment: Winter Cooling: Brumation: Hibernation: northerly latitudes. Aestivation: Photoperiod (as usually given in captivity) Photoperiod - 14:10h summer:winter Temperate - 13:11h summer:winter; Subtropical - 12h all year; Tropical Microhabitat wetlands; J - Aquatic. I - Riparian or H - Arboreal; G - Semi-arboreal; savanna; Grassland or F - or shrubs; Foliage E - or burrows; crevices Rocks, floor; D - Forest C - litter; B - Leaf A - Fossorial; Scientific name Crocodilia Caiman crocodilus Rhynchocephalia Sphenodon punctatus Crocodylus mindorensis Crocodylus Crocodylus moreletii Crocodylus Osteolaemus tetraspis Paleosuchus palpebrosus

Journal of Zoo and Aquarium Research 4(1) 2016 53 Baines et al. E G G H H H H H H H D D DF DF DF EG EG EG EH EH EH GH CEI EGH DEG ABD DEFG Microhabitat 10 20 20 14–18 14–18 10–15 18–22 23–25 21–23 20–22 15–24 20–23 20–22 22–25 22–25 22–25 18–22; 5 different) Winter (if Winter (Brumation) temperature (°C) 21 21 Night – ambient (air) 18–22 18–22 20–25 20–25 20–24 20–25 20–25 24–26 20–25 20–24 15–20 15–20 25–27 24–26 23–25 22–25 15–26 23–25 20–26 24–28 24–28 23–25 25–28 25–28 Summer 28 26 26 18–22 18–22 18–20 24–26 17–27 24–26 22–26 20–30 25–27 26–30 30–32 28–32 28–32 different) Winter (if Winter (Brumation) 25–30; 10–15 25–30; 30 28 28 temperature (°C) Day – ambient (air) 23–28 23–28 25–30 25–30 25–27 25–30 25–30 25–30 27–32 26–30 21–26 20–30 25–37 25–35 26–28 25–30 20–30 27–29 25–32 30–35 30–35 25–33 30–35 30–35 Summer 30 32 40 40 40 40 30–32 30–32 30–35 30–35 30–40 34–36 25–30 30–35 30–32 30–32 29–32 35–40 35–45 30–35 30–35 40–48 40–50 40–45 40–45 40–50 40–50 Basking zone – Substrate surface temperature (°C) (if any) Cooling Cooling Cooling Cooling Cooling Hibernation Brumation / Winter treatment Winter 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 14:10 12:12 12:12 12:12 12:12 12:12 12:12 12:12 14:10 12:12 Photoperiod 12:12/13:11 12:12/13:11 13:11/14:10 1 2 1 2 1 2 2 3 1 3 3 2 3 2 3 2 3 2 4 3 4 4 3 3 1–2 3–4 3–4 Zone Ferguson Ferguson 1 4 1 1 1 1 1 1 1 1 2 2 1 1 7 7 13 14 13 10/13 08/13 02/07 02/07 04/05/ Biome Common Name Rough-bellied Mountain Horned Dragon Anole American Green Two-horned Mountain Horned Two-horned Dragon Jamaican Blue-pants Anole Jamaican Blue-pants Jamaican Brown Anole Jamaican Brown Plumed Basilisk Martinique Anole Martinique Anole Cuban Brown Fiji Banded Iguana Brown Leaf Chameleon Bornean bloodsucker/Green Crested Lizard Oriental Garden Lizard, Eastern Garden Lizard, Bloodsucker Four Horned Chameleon Frilled Lizard Meller's Chameleon Yemen Chameleon Yemen Giant Hispaniolan Galliwasp Parson's Chameleon Prehensile or Monkey-tailed Skink Collared Lizard Utila Iguana Guatemalan Black Iguana Central Netted Dragon Rhinoceros Iguana Cuban Rock Iguana Cayman Brac Iguana/ Sister Isles Iguana Grand Cayman Iguana/ Blue Iguana Scientific name Acanthosaura lepidogaster Anolis carolinensis Squamata: Lacertilia Acanthosaura capra Anolis grahami Anolis lineatopus Basiliscus plumifrons Basiliscus plumifrons Anolis roquet summus Anolis roquet Anolis sagrei Brachylophus bulabula Brookesia superciliaris Brookesia Bronchocela cristatella Bronchocela Calotes versicolor Chameleo trioceros Chameleo trioceros quadricornis Chlamydosaurus kingii Chamaeleo melleri Chamaeleo calyptratus Celestus warreni Calumma parsonii Corucia zebrata Crotaphytus collaris Crotaphytus Ctenosaura bakeri Ctenosaura palearis Ctenophorus nuchalis Cyclura cornuta Cyclura nubila Cyclura nubila caymanensis Cyclura nubila lewisi

54 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians I I F A D D D D H H H H D G H DF DF EH EH DE DE EH BD AG GH DFG ABCEF BDEFG Microhabitat 20 20 2–6 5–15 9–10 5–15 10–15 10–15 10–15 18–24 16–20 20–22 10–15 23–25 10–15 20–22 20–24 18–22 18–20 different) Winter (if Winter 18–20; 4–8 (Brumation) temperature (°C) 25 20 Night – ambient (air) temps 20–25 24–26 20–24 20–24 20–25 18–24 20–24 22–24 22–24 24–26 20–25 20–25 25–28 20–25 24–26 10–20 23–25 24–26 20–25 22–24 20–22 22–24 16–22 Summer ambient UK 2–6 5–15 15–25 15–20 20–25 24–28 20–24 10–20 26–28 10–12 20–25 10–15 28–32 20–25 27–30 24–28 24–28 22–24 20–23 25–27 24–28; 12 – 15 different) Winter (if Winter (Brumation) 30 temperature (°C) Day – ambient (air) temps 25–35 25–28 28–32 25–29 25–30 25–30 26–30 25–30 26–29 26–29 28–32 24–30 25–30 25–30 30–35 25–30 30–32 25–35 30–35 27–32 25–30 26–30 26–28 24–27 28–30 24–28 Summer ambient UK 50 32 40 35 32 32 35 35 35 None 30–35 35–40 35–40 30–32 35–40 35–40 34–37 30–40 40–50 35–40 30–40 40–50 35–40 35–45 30–32 28–32 30 –35 ambient UK temps Basking zone – Substrate surface temperature (°C) (if any) Cooling Cooling Cooling Cooling Cooling Cooling Cooling/ Brumation Brumation Brumation Brumation Brumation Hibernation None / Cooling Winter treatment Winter 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 14:10 14:10 12:12 12:12 12:12 12:12 14:10 14:10 14:10 14:10 14:10 12:12 12:12 12:12 14:10 12:12 14:10 12:12/13:11 12:12/13:11 Photoperiod 3 2 1 3 3 3 1 2 2 2 2 2 4 4 2 3 4 3 4 2 3 1 1 2 3–4 2–3 3–4 2–3 Zone Ferguson Ferguson 1 4 7 1 2 1 1 8 2 1 4 1 7 1 2 7 7 7 13 13 13 13 13 01/02 01/02 04/13 Biome 01/04/07 08/04/12 Common Name Northern Caiman Lizard Desert Iguana Cunningham's Rock Skink Leopard Gecko Berber Skink Panther Chameleon Sudan Plated Lizard Tokay Gecko Tokay Angle-headed Dragon Rio Fuerte Beaded Lizard Angle-headed Dragon Bell's Forest Dragon Gila Monster Skink Pink-Tongued Lesser Earless Lizard Antillean Iguana Lesser Australian Water Dragon Water Australian Sand Lizard Serrated Casque-headed Iguana Painted Dragon (Starred Agama spp) Painted Dragon (Starred Curly Tail Lizard Tail Curly Round Island Skink Fire Skink Striped Water Dragon Water Striped Electric blue day gecko / Turquoise Turquoise Electric blue day gecko / dwarf gecko Lesser Night Gecko Northern Velvet Gecko Velvet Northern Scheltopusik Scientific name Dracaena guianensis Dipsosaurus dorsalis Egernia cunninghami Eublepharis macularius Eumeces schneideri pardalis Furcifer Gerrhosaurus major Gerrhosaurus Gecko gecko Gonocephalus grandis Heloderma horridum exasperatum Gonocephalus doriae Gonocephalus bellii Heloderma suspectum Hemisphaeriodon gerrardii Holbrookia maculata Holbrookia Iguana delicatissima Intellagama (Physignathus) lesueurii Lacerta agilis Laemanctus serratus Laudakia stellio brachydactyla Leiocephalus carinatus Leiolopisma telfairi Lepidothyris (Riopa) fernandi Lophognathus temporalis Lygodactylus williamsi Lygodactylus Nactus coindemirensis Oeudura castelnaui Ophisaurus apodus

Journal of Zoo and Aquarium Research 4(1) 2016 55 Baines et al. D D D H D H H EF CF DF EH DE EG EH EH DG EGI DEF EFG BEH EGH EGH ABCI ABCF BCDF Microhabitat 20 5–10 5–10 5–10 14–20 18–22 18–20 10–15 20–22 10–15 10–15 15–20 16–20 18–20 10–15 22–25 different) Winter (if Winter (Brumation) 20–22;10–15 temperature (°C) 20 22 28 Night – ambient (air) 11–16 20–24 23–25 16–17 20–24 18–22 20–25 15–20 24–26 20–25 15–20 20–25 25–30 18–20 23–25 20–25 20–25 20–22 20–25 25–30 23–25 23–25 Summer 20 5–20 5–10 18–28 24–28 22–28 15–20 10–15 24–28 10–15 10–15 25–30 19–23 22–24 10–15 25–28 different) Winter (if Winter (Brumation) 25–30; 15–20 temperature (°C) 27 28 Day – ambient (air) 23–28 24–25 28–32 25–30 28–32 26–30 25–30 20–30 28–35 27–33 25–30 25–30 18–20 24–30 30–35 25–30 25–28 25–29 26–28 30–35 30–35 25–30 25–30 Summer 36 35 35 25 50 50 28 33 29 28–32 35–40 35–45 35–40 35–40 30–32 35–40 35–40 30–40 30–40 40–45 30–40 30–40 35–45 30–35 30–35 Basking zone – Substrate surface temperature (°C) (if any) Cooling Cooling Cooling Cooling Cooling Cooling Cooling Cooling/ Cooling/ Brumation Brumation Brumation Brumation Brumation Brumation Brumation Brumation Brumation Hibernation Brumation / Winter treatment Winter 13:11 13:11 13:11 13:11 13:11 12:12 12:12 14:10 14:10 14:10 14:10 14:10 14:10 14:10 14:10 12:12 14:10 12:12 12:12 14:10 14:10 14:10 Photoperiod 13:11/14:10 13:11/14:10 13:11/14:10 1 3 2 2 4 4 3 4 3 2 4 4 2 1 4 3 3 3 2–3 2–3 2–3 2–3 3–4 2–3 2–3 Zone Ferguson Ferguson 1 1 4 7 8 4 1 1 1 1 2 1 2 13 12 10 13 13 07/08 08/12 12/13 05/13 01/10 12/13 08/13 Biome 04/07/ 04/08/ 07/08/ 01/02/04/ Common Name Crocodile Skink chameleon Jackson’s Black-and-White Tegu Black-and-White Eastern Blue-tongued Lizard Shingleback Lizard Southern or Blotched Blue-tongued Lizard Wonder Gecko Wonder Moorish Gecko Sungazer, Giant Girdled Lizard Sungazer, Blue Spiny Lizard Texas spiny lizard Texas Sagebrush Lizard Eastern Fence Lizard Bearded Pygmy Chameleon Chuckwalla Angel Island Chuckwalla Inland Bearded dragon Spiny-headed Tree Lizard Tree Spiny-headed Gargoyle Gecko Gargoyle Crested Gecko Asian Water Dragon Water Asian Texas Horned Lizard Texas Yellow Headed Day Gecko Yellow Giant Day Gecko Standing's Day Gecko Scientific name Tribolonotus gracilis Tribolonotus jacksonii Trioceros Tupinambis merianae Tupinambis Tiliqua scincoides Tiliqua Tiliqua rugosa Tiliqua Tiliqua nigrolutea Tiliqua Teratoscincus scincus Teratoscincus Tarentola mauritanica Tarentola Smaug (Cordylus) giganteus Smaug (Cordylus) Sceloporus serrifer cyanogenys Sceloporus olivaceus Sceloporus graciosus Sceloporus consobrinus (Louisiana USA) Rieppeleon brevicaudatus Sauromalus ater Sauromalus hispidus Sauromalus Pogona vitticeps Plica plica Rhacodactylus auriculatus Rhacodactylus ciliatus Physignathus cocincinus Phrynosoma cornutum Phelsuma klemmeri Phelsuma madagascariensis grandis Phelsuma standingi

56 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians C G H H H H G DF DF DF DF FG DF BC BC EH EH DE AD CGI CFG ADF CDE BEG ADFI CDFG BCEG Microhabitat 20 23 22–25 20–25 20–24 20–24 23–24 23–24 16–22 22–24 20–22 18–20 10–18 17–19 15–18 10–15 21–23 18–23 25–28 22–25 22–25 18–20 21–23 different) Winter (if Winter temperature (°C) 22 20 23 Night – ambient (air) 24–28 25–28 25–28 25–28 28–29 28–29 18–24 24–26 24–26 20–25 16–18 18–20 18–20 20–25 23–26 24–25 20–24 24–26 24–26 24–26 22–26 25–28 23–26 Summer 30 28 25–30 25–30 25–28 25–28 25–26 25–26 20–26 24–26 25–27 25–30 20–25 21–23 16–20 18–20 28–30 28–35 22–25 30–34 26–30 26–30 28–30 25–27 different) Winter (if Winter 26 30 temperature (°C) Day – ambient (air) 30–35 28–35 28–30 28–30 29–30 29–30 24–30 26–28 27–30 30–38 28–35 23–25 20–25 25–30 28–35 26–30 30–40 25–30 30–32 28–32 28–32 26–32 30–32 30–35 28–30 Summer 32 32 32 45 40 None 40–45 40–45 28–30 30–35 30–35 45–50 45–50 40–50 25–28 30–40 40–50 31–36 55–65 35–40 35–40 35–40 35–40 40–50 34–36 Basking zone – Substrate surface temperature (°C) (if any) Cooling Cooling Cooling Cooling Cooling Cooling Cooling Cooling Winter treatment Winter Cooling Cooling Cooling Cooling Cooling Cooling Cooling 13:11 13:11 14:10 12:12 14:10 12:12 14:10 14:10 12:12 12:12 12:12 12:12 12:12 12:12 12:12 12:12 12:12 13:11 13:11 13:11 13:11 13:11 12:12 12:12 Photoperiod 12:12/13:11 13:11/14:10 13:11/14:10 4 4 4 1 3 3 3 3 4 2 2 2 3 3 2 1 1 1 2 2 1 1 1–2 3–4 3–4 1–2 1–2 Zone Ferguson Ferguson 1 1 7 7 1 1 1 7 2 2 7 1 1 1 1 1 13 13 13 13 13 13 01/02 01/14 01/02 Biome 04/07/08 07/09 Common Name Dumeril's Boa Children's Python Egyptian Uromastyx / Mastigure/ Dab Lizard Stimson's Python Saharan Uromastyx / Spinytailed Lizard Woma python Woma Gaboon Viper Gaboon Ornate Uromastyx Rhinoceros Viper Rhinoceros Boa Constrictor Henkel’s leaf-tail gecko Henkel’s Banded Mangrove Snake Satanic Leaf Tailed Gecko Tailed Satanic Leaf Eyelash Viper Eyelash Desert Side-blotched Lizard Solomon Island Boa Black Tree Monitor Tree Black Philippine Water Monitor Water Philippine Bosc Monitor, Savannah Monitor Bosc Monitor, Kimberley Rock Monitor Komodo Dragon Blue Tree Monitor Tree Blue Emerald Tree Monitor Tree Emerald Crocodile Tree Monitor Tree Crocodile Spencer’s goanna Spencer’s Timor Monitor Timor Lace Monitor Scientific name Squamata: Serpentes Squamata: Serpentes Acrantophis dumerili Antaresia childreni childreni Antaresia Uromastyx aegyptia Uromastyx Antaresia stimsoni orientalis Antaresia Uromastyx geyri Uromastyx Aspidites ramsayi Bitis gabonica Uromastyx ornata Uromastyx Bitis nasicornis Boa constrictor Uroplatus henkeli Uroplatus Boiga dendrophila melanota Boiga dendrophila Uroplatus phantasticus Uroplatus Bothriechis schlegelii Uta stansburiana stejnegeri Candoia carinata Varanus beccarii Varanus Varanus cumingi Varanus Varanus exanthematicus Varanus Varanus glauerti Varanus Varanus komodoensis Varanus Varanus macraei Varanus Varanus prasinus Varanus Varanus salvadorii Varanus Varanus spenceri Varanus Varanus timorensis Varanus Varanus varius Varanus

Journal of Zoo and Aquarium Research 4(1) 2016 57 Baines et al. I E C H H IJ H D G H D EF DF CF DF EH DF CG GH EHI CGI EFG BCD ADEF DEFG DEFG DEGH BCDEGH Microhabitat 16 9–12 20–22 20–22 22–24 20–24 14–16 10–15 10–15 18–21 15–20 22–26 15–20 10–12 12–17 22–24 different) Winter (if Winter temperature (°C) 20 Night – ambient (air) 24–26 24–26 24–26 24–26 22–26 25–27 20–24 24–26 24–26 20–23 24–26 18–22 24–26 26–28 20–26 24–26 18–20 18–20 17–19 22–26 22–26 24–26 22–26 Summer 19 20–25 20–25 25–27 26–28 24–28 14–18 10–15 10–15 22–25 20–24 24–28 22–26 25–27 10–12 12–17 17–19 27–29 different) Winter (if Winter temperature (°C) Day – ambient (air) 25–30 25–30 26–30 28–30 28–30 28–32 28–32 26–32 28–32 24–30 24–28 24–28 24–28 25–30 24–28 26–30 28–30 25–28 28–30 26–30 18–32 22–25 24–26 23–25 28–32 28–32 28–30 28–32 Summer 32 30 31 30–35 30–35 30–32 30–32 35–40 30–45 35–40 28–30 28–32 28–32 30–35 30–35 30–32 30–32 34–36 22–28 35–40 Basking zone – Substrate surface temperature (°C) (if any) Winter treatment Winter Night cooling Cooling Cooling / Brumation Brumation Brumation Cooling Cooling Cooling Cooling Cooling Brumation Brumation Cooling 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 12:12 12:12 12:12 12:12 12:12 13:11 14:10 13:11 13:11 13:11 13:11 12:12 12:12 14:10 12:12 12:12 14:10 14:10 12:12 13:11/14:10 Photoperiod 2 1 1 2 2 2 1 1 1 2 2 2 3 1 1 1 1 2 1 1–2 1–2 1–2 1–2 1–2 1–2 1–2 1–2 1–2 Zone Ferguson Ferguson 1 1 1 7 2 1 1 2 1 1 4 7 1 1 7 1 2 1 13 13 01/07 08/10 02/13 07/02 01/14 04/08 01/02 Biome 01/04/05 Common Name Horned Viper Horned Emerald Tree Boa Tree Emerald Viper White Lipped Green Mamba Black Mamba Cuban Boa Jamaican Boa Green Anaconda Green Red–tailed Ratsnake Western Hognose Snake Western Pueblan Milksnake Sinaloan Milksnake Stuart's Milksnake Savu Python Amethystine Python Boelens Python Central Carpet Python/ Bredl's Python Diamond Python Top End Carpet Python Top Rough Green Snake Green Tree Python Tree Green Hundred Flower Snake Corn Snake Mang Mountain Viper Mang Mountain Borneo Short–tailed Python Burmese Python Royal Python Reticulated Python Scientific name Cerastes cerastes Corallus caninus albolabris Cryptelytrops Dendroaspis angusticeps Dendroaspis Dendroaspis polylepis Dendroaspis Epicrates angulifer Epicrates subflavus Eunectes murinus Gonyosoma oxycephalum Heterodon nasicus Heterodon Lampropeltis triangulum Lampropeltis campbelli Lampropeltis triangulum Lampropeltis sinaloae Lampropeltis triangulum Lampropeltis stuarti Liasis macklotti savuensis Morelia amethistina Morelia Morelia boeleni Morelia Morelia bredli bredli Morelia Morelia spilota Morelia Morelia spilota variegata Morelia Opheodrys aestivus Morelia viridis Morelia Orthriophis moellendorffi Pantherophis guttatus Pantherophis guttatus Protobothrops mangshanensis Protobothrops Python brietensteini Python molurus bivittatus Python regius Python (Broghammerus) Python (Broghammerus) reticulatus

58 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians I E G G H H H H H H DF EG EG EH EH EG EH GH CEI EGH EGH DEG ABD EGHI BCDF DEFG Microhabitat 10 2–6 14–18 14–18 10–15 15–20 21–23 15–24 20–22 18–22 22–25 14–18 23–25 20–23 20–22 22–25 18–22; 5 different) Winter (if Winter (Brumation) temperature (°C) 20 22 18 Night – ambient (air) 22–24 22–24 24–28 24–28 21–25 22–28 22–24 25–28 24–28 22–24 20–24 10–16 18–24 22–24 22–24 24–26 20–25 Summer 3–7 3–7 3–7 3–7 2–6 3–7 5–10 3–10 24–26 24–28 24–26 26–30 20–25 20–28 22–24 24–28 28–30 25–30 24–26 10–15 16–20 22–24 24–28 22–25 25–27 different) Winter (if Winter 25–30 22–30 26–28 28–30 28–32 22–30 28–32 25–29 20–30 25–31 22–30 20–25 25–28 28–30 30–35 30–35 26–28 26–30 18–24 22–28 24–26 28–30 25–30 20–25 29–31 27–30 temperature (°C) Day – ambient (air) Summer 35 35 35 35 35 35 35 35 30 35 30 35 30 35–40 35–45 35–50 30–35 30–34 28–32 45–50 40–45 30–35 28–30 30–35 35–45 30–35 Basking zone – Substrate surface temperature (°C) (if any) Cooling Cooling Cooling Cooling Cooling Cooling Cooling Cooling Cooling Cooling/ Cooling/ Brumation Brumation Brumation Brumation Hibernation Hibernation Hibernation Hibernation Hibernation Hibernation Hibernation Cooling/ None Winter treatment Winter 13:11 13:11 13:11 13:11 14:10 14:10 12:12 12:12 14:10 14:10 14:10 12:12 14:10 14:10 12:12 14:10 12:12 14:10 12:12 14:10 12:12 12:12 Photoperiod 13:11/14:10 13:11/14:10 13:11/14:10 13:11/14:10 3 2 3 3 3 2 3 2 3 3 2 3 3 3 2–3 2–3 3–4 2–3 1–2 3–4 1–2 2–3 2–3 2–3 1–2 2–3 Zone Ferguson Ferguson 4 1 1 7 4 7 1 1 4 1 4 4 1 1 4 1 4 13 13 08/10 08/12 08/12 06/08 01/07 07/13 04/07/ 04/07/ 04/05/ 01/02/ Biome 04/08/12 Common Name Smooth Softshell Rhinoceros Rat Snake Tortoise, Russian Horsfield's Tortoise Roti Island snake–necked turtle Sulcata Tortoise, African Spurred Tortoise, Sulcata Tortoise Ploughshare Tortoise Ploughshare Spiny Softshell Tortoise Radiated Madagascan Tree Boa Tree Madagascan Vietnamese Keeled Turtle Keeled Vietnamese Painted Turtle Painted Turtle Flowerback Box Vietnamese/ Common Snapper Turtle Spotted Yellow footed Tortoise footed Yellow Common or Eastern snake–necked turtle Golden coin Box Turtle Golden coin Box Broad– shelled Turtle Broad– shelled Adder San Fransico Garter Snake Zhou's Box Turtle Zhou's Box Murray Short–necked Turtle Murray Short–necked Turtle European Pond Aldabran Tortoise Aldabran Red Foot Tortoise Red Foot Indian Star Tortoise Indian Star Scientific name Apalone mutica Rhyncophis boulengeri Chelonia Agrionemys horsfieldii Chelodina mccordi Centrochelys (Geochelone) Centrochelys sulcata Astrochelys yniphora Astrochelys Apalone spinifera radiata Astrochelys Sanzinia madagascarensis Cuora mouhotti Crysemys picta ssp. Cuora galbinifrons Chelydra serpentina Clemmys guttata Chelonoidis denticulata Chelodina longicollis Cuora trifasciata Chelodina expansa Vipera berus Vipera Thamnophis sirtalis tetrataenia Cuora zhoui Emydura macquarii Emys orbicularis Geochelone gigantea/ Dipsochelys dussumieri Geochelone carbonaria Geochelone elegans

Journal of Zoo and Aquarium Research 4(1) 2016 59 Baines et al. I I F A H D D D D D D H H H H D G DF DF DF DF EH EH DE DE EH AG GH DFG BDEFG Microhabitat 20 20 20 20 5–15 9–10 5–15 22–25 22–25 10–15 10–15 10–15 18–24 16–20 20–22 10–15 23–25 10–15 20–22 20–24 18–22 different) Winter (if Winter 18–20; 4–8 (Brumation) 22 temperature (°C) Night – ambient (air) 24–28 24–28 22–24 24–26 20–25 20–25 22–25 22–24 23–25 23–25 22–24 24–26 Summer 14 12 14 15 3–7 3–7 3–7 3–7 3–7 3–7 3–7 3–7 3–10 3–10 3–10 3–10 26–30 26–30 22–24 24–26 20–25 22–24 24–26 23–26 24–26 24–26 24–26 different) Winter (if Winter temperature (°C) Day – ambient (air) 24–26 22–30 24–30 28–32 28–32 25–30 25–30 25–30 25–30 26–28 22–30 25–30 28–30 25–30 26–28 25–30 25–30 26–28 25–30 25–30 25–30 25–30 25–30 25–30 28–32 22–30 22–30 22–30 22–30 22–30 Summer 30 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35 35–45 35–45 40–50 28–32 30–32 30–35 30–35 26–28 35–40 Basking zone – Substrate surface temperature (°C) (if any) Cooling Cooling Cooling Cooling Cooling Cooling Cooling Brumation Brumation Brumation Brumation Hibernation Hibernation Hibernation Hibernation Hibernation Hibernation Hibernation Hibernation Hibernation Hibernation Hibernation Hibernation Winter treatment Winter 13:11 13:11 13:11 13:11 13:11 13:11 14:10 12:12 14:10 12:12 12:12 12:12 14:10 14:10 14:10 12:12 12:12 14:10 12:12 12:12 12:12 14:10 14:10 14:10 14:10 14:10 14:10 13:11/14:10 13:11/14:10 13:11/14:10 Photoperiod 1 2 2 4 3 1 3 3 3 3 3 2 2 2 2 3–4 3–4 1–2 2–3 2–3 2–3 2–3 3–4 3–4 2–3 3–4 3–4 2–3 2–3 2–3 Zone Ferguson Ferguson 1 4 1 7 7 4 4 1 4 4 4 7 1 4 4 1 4 5 4 5 1 2 4 4 4 4 4 02/07 01/07 01/02 Biome Common Name Black–breasted Leaf Turtle Black–breasted Leaf Turtle Wood Tortoise Elongated Galapagos Tortoise Galapagos Tortoise Leopard Turtle Ouachita Map False, Mississippi, Northern Map Turtle Turtle Spiny Bell's Hingeback Tortoise Bell's Hingeback Home's Hingeback Tortoise Home's Hingeback Turtle Eastern Mud Diamond Back Terrapin Diamond Back Turtle Mediterranean Pond Pancake Tortoise Pancake Annam Leaf Turtle Annam Leaf Turtle Reeves Turtle Eurasian Pond Yellow–spotted Amazon River Turtle River Amazon Yellow–spotted Cooters Florida Red Bellied Cooter Malaysian Giant Pond Turtle Malaysian Giant Pond Turtle Side–neck Red Bellied Cooter Turtle Wood Painted Flat–tailed Tortoise Flat–tailed Razorback Musk Loggerhead Musk Common Musk Turtle Eastern Box Ornate Box Turtle Ornate Box Scientific name Geoemyda spengleri Glyptemys insculpulata Indotestudo elongata Geochelone pardalis Graptemys ouachitensis Graptemys pseudogeographica ssp Heosemys spinosa Geochelone nigra Kinixys belliana Kinixys homeana Kinosternon subrubrum Malaclemys terrapin leprosa Mauremys Malacochersus tornieri Mauremys (Annamemys) Mauremys annamensis reevesii Mauremys rivulata Mauremys Podocnemis unifilis Pseudemys concina ssp. Pseudemys nelsoni Orlitia borneensis Phrynops geoffranus Pseudemys rubriventris Rhinoclemmys pulcherrima Pyxis planicauda Sternotherus carinatus Sternotherus minor Sternotherus odoratus carolina Terapene Terapene ornata Terapene

60 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians I H H H H H G H CI DI EF EH EH BD BCI BDI BCG BCD BDFI DEFIJ DEFIJ BCEG BCEG BDEGI ABCEF Microhabitat 11 2–6 8–12 5–10 18–20 14–16 17–18 22–25 10–12 10–15 18–20 22–24 22–24 20–22 20–22 20–22 20–22 20–22 20–22 different) Winter (if Winter 18 (spring) (winter); 16– 2–8 (winter); 2–8 (winter); 6–12 (spring) 6–12 (spring) 22 17 temperature (°C) Night – ambient (air) 11–14 11–14 10–15 10–15 17–20 18–19 22–25 20–24 24–26 13–15 18–14 16–18 20–23 22–26 24–26 20–24 20–24 20–24 20–24 20–23 22–24 21–23 Summer 17 4–7 4–7 2–6 3–10 3–10 8–20 5–10 15–22 22–23 15–17 22–25 24–26 24–28 22–25 22–25 22–25 22–25 22–25 22–24 23–25 different) Winter (if Winter 25 (spring) (winter); 22– 3–8 (winter); 3–8 (winter); 8–21 (spring) 8–21 (spring) 22 temperature (°C) Day – ambient (air) 20–30 22–26 22–30 23–24 25–30 25–30 28–32 28–30 25–30 18–20 24–28 23–25 18–26 24–26 18–23 26–30 28–32 24–28 24–28 24–28 24–28 24–26 24–28 24–26 Summer 35 35 30 35 35 35 25 26 30–35 35–45 25–30 30–35 30–32 Basking zone – Substrate surface temperature (°C) (if any) Cooling Cooling Cooling Cooling Cooling Cooling Cooling Cooling Dry spell Brumation Brumation Hibernation Hibernation Hibernation Hibernation Winter treatment Winter 13:11 13:11 13:11 13:11 13:11 13:11 14:10 14:10 14:10 14:10 14:10 12:12 14:10 14:10 14:10 14:10 14:10 12:12 12:12 12:12 12:12 12:12 12:12 12:12 Photoperiod 13:11/14:10 3 3 1 3 4 3 1 1 1 1 2 2 1 1 1 1–2 3–4 3–4 3–4 1–2 1–2 1–2 1–2 1–2 1–2 Zone Ferguson Ferguson 1 1 4 7 1 1 1 1 1 1 1 1 12 12 12 12 03/14 04/08 06/12 01/07 01/02 01/02/ 04/05/ Biome 04/05/08 04/05/12 04/05/09 01/02/14 Common Name Spur–thighed Tortoise Spur–thighed Tortoise Hermann's Frog Tree Red-eyed Lemur Leaf Frog Frog Tree Black-eyed Yellow Bellied Slider Yellow Marginated Tortoise Marginated Egyptian Tortoise Egyptian Hispaniolan Elegant Slider Red–eared Slider Toad Mallorcan Midwife Common Midwife Toad Common Midwife Yellow-bellied Toad Yellow-bellied Harlequin Toad Harlequin Oriental or Chinese Fire-bellied Toad Toad Common Bony-headed Toad Bony-headed Cane Toad Cane Splendid Leaf Frog Bumblebee Dart Frog Dyeing Dart Frog Blue Dart Frog Amazonian Dart Frog Frog Tomato Sambava Green and Black Dart Frog Scientific name Testudo hermanni Testudo Agalychnis lemur Agalychnis moreletii Testudo graeca ibera/ Testudo graeca ibera/ Testudo Testudo ibera Anura Agalychnis callidryas Trachemys scripta Trachemys Testudo marginata Testudo Testudo kleinmanni Testudo decorata Trachemys scripta elegans Trachemys Alytes muletensis Alytes obstetricans Bombina variegata Atelopus spumarius hoogmoedi Bombina orientalis Bufo bufo Bufo galeatus Bufo marinus Cruziohyla calcarifer Dendrobates leucomelas Dendrobates tinctorius Dendrobates tinctorius / Dendrobates azureus ventrimaculatus Dendrobates Dyscophus guineti Dendrobates auratus Dendrobates

Journal of Zoo and Aquarium Research 4(1) 2016 61 Baines et al. H G G H HI EH BC BC BD BD BCI BDI BDI BDI BDE BCH BEFG DEFIJ BCEG BCDE DEFHI BCEGH Microhabitat 17 22–24 18–21 18–21 16–18 18–20 16–20 22–24 10–18 18–20 16–17 18–20 18–20 22–24 different) Winter (if Winter 2–8 (winter); 6–12 (spring) 17 20 temperature (°C) Night – ambient (air) 24–26 18–21 18–22 11–14 22–24 18–20 20–22 22–24 22–24 22–26 22–24 20–22 20–22 17–19 20–22 20–24 24–26 22–24 20–24 18 –24 Summer 22 26–30 20–24 20–24 22–24 18–20 22–25 22–24 22–24 20–25 24–26 22–24 18–22 22–24 20–24 22–24 23–25 24–26 22–24 20–24 different) Winter (if Winter 3–8 (winter); 8–21 (spring) 22 25 temperature (°C) Day – ambient (air) 28–32 20–24 20–26 24–28 20–22 24–28 26–28 26–28 22–28 26–30 24–28 24–26 24–26 24–27 24–26 26–28 25–27 18–23 24–28 26–30 Summer 25 36 30 26 32–40 30–35 30–35 30–35 Basking zone – Substrate surface temperature (°C) (if any) Cooling Cooling Cooling Cooling Cooling Cooling Winter treatment Winter 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 13:11 12:12 12:12 12:12 12:12 14:10 14:10 14:10 14:10 14:10 Photoperiod 1 3 1 2 2 2 2 1 2 2 3 2 2 1 2 2 2 1 2 2 2 1–2 Zone Ferguson Ferguson 1 1 1 1 1 1 1 1 1 1 7 1 1 1 1 2 1 01/02 08/09 01/02 01/07 02/07/ Biome 04/05/12 Common Name Amazon Milk Frog Bug-eyed Tree Frog Tree Bug-eyed Vietnamese Mossy Frog Vietnamese Feae’s Flying Frog Feae’s Reticulated Poison Frog Pasco Poison Frog Golden Tree Frog Tree Golden Golfodulcean Poison Frog Golden Poison Frog Black-legged Dart Frog White's Tree Frog Tree White's Strawberry Poison Frog Toad Tree Borneo Morogoro Tree Toad Tree Morogoro Long-nosed Horned Frog Green Mantella Frog Golden Mantella Frog Green-eyed Frog Natterjack Toad Natterjack Maranon Poison Frog Mountain Chicken Phantasmal Poison Frog Scientific name Trachycephalus resinifictrix Trachycephalus Theloderma stellatum Theloderma corticale Rhacophorus feae Ranitomeya reticulata Ranitomeya lamasi Polypedates leucomystax Phyllobates vittatus Phyllobates terribilis Phyllobates bicolor Pelodryas caerulea Oophaga pumilio Pedostibes hosii Nectophrynoides viviparus Megophrys nasuta Mantella viridis Mantella aurantiaca Lithobates vibicarius Epidalea (Bufo) calamita Excidobates mysteriosus Leptodactylus fallax Epipedobates anthonyi

62 Journal of Zoo and Aquarium Research 4(1) 2016 A UV-B lighting guide for reptiles and amphibians I IJ H H H CI DI BDI Microhabitat 11 8–12 18–20 27–28 17–18 10–12 26–28 different) Winter (if Winter 16–18 (spring) 14–16 (winter); temperature (°C) Night – ambient (air) 11–14 11–14 11–14 25–28 28–30 18–20 22–25 28–30 Summer 8 –21 15–18 27–28 26–28 Oct-15 Dec-14 (spring) different) Winter (if Winter 3–8 (winter); 3–8 (winter); 3–8 (winter); 8–21 (spring) 8–21 (spring) temperature (°C) Day – ambient (air) 25–28 18–23 18–23 18–23 28–30 18–20 24–30 28–30 Summer 30–35 Basking zone – Substrate surface temperature (°C) (if any) Cooling Cooling Cooling Cooling Cooling Cooling Winter treatment Winter 13:11 14:10 14:10 14:10 12:12 14:10 14:10 12:12 Photoperiod 1 1 1 1 1 1 2 1–2 Zone Ferguson Ferguson 1 1 12 03/10 05/08 10/12 08/12 05/08 06/04/ 04/05/ 06/04/ Biome Himalayan Newt Great Crested Newt Kaisers Newt Fire Salamander Aquatic Caecilian, Rubber Eel, Worm. Caecilian Smooth or Common Newt Sardinian Brook Salamander Rio Cauca Caecilian Common Name Scientific name Tylototriton verrucosus Tylototriton Triturus cristatus Triturus Neurergus kaiseri Neurergus Salamandra salamandra Typhlonectes spp. Typhlonectes Lissotriton (Triturus) vulgaris Lissotriton (Triturus) Caudata platycephalus Euproctus Gymophiona: Caeciliidae natans Typhlonectes

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